Ljerka Ostojić

The Evolution of Self-Control

Significance Although scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problem-solving tasks measuring self-control and evaluated the leading hypotheses regarding how and why cognition evolves. Across species, differences in absolute (not relative) brain volume best predicted performance on these tasks. Within primates, dietary breadth also predicted cognitive performance, whereas social group size did not. These results suggest that increases in absolute brain size provided the biological foundation for evolutionary increases in self-control, and implicate species differences in feeding ecology as a potential selective pressure favoring these skills. Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.

Behavioural coordination of dogs in a cooperative problem-solving task with a conspecific and a human partner

The process of domestication has arguably provided dogs (Canis familiaris) with decreased emotional reactivity (reduced fear and aggression) and increased socio-cognitive skills adaptive for living with humans. It has been suggested that dogs are uniquely equipped with abilities that have been identified as crucial in cooperative problem-solving, namely social tolerance and the ability to attend to other individuals’ behaviour. Accordingly, dogs might be hypothesised to perform well in tasks in which they have to work together with a human partner. Recently, researchers have found that dogs successfully solved a simple cooperative task with another dog. Due to the simplicity of the task, this study was, however, unable to provide clear evidence as to whether the dogs’ successful performance was based on the cognitive ability of behavioural coordination, namely the capacity to link task requirements to the necessity of adjusting one’s actions to the partner’s behaviour. Here, we tested dogs with the most commonly used cooperative task, appropriate to test behavioural coordination. In addition, we paired dogs with both a conspecific and a human partner. Although dogs had difficulties in inhibiting the necessary action when required to wait for their partner, they successfully attended to the two cues that predicted a successful outcome, namely their partner’s behaviour and the incremental movement of rewards towards themselves. This behavioural coordination was shown with both a conspecific and a human partner, in line with the recent findings suggesting that dogs exhibit highly developed socio-cognitive skills in interactions with both humans and other dogs.

Dominance, pair bonds and boldness determine social-foraging tactics in rooks, Corvus frugilegus

Socially foraging animals can search for resources themselves (produce) or exploit the discoveries made by others (scrounge). The extensive literature on producerescrounger dynamics has mainly focused on scramble competition over readily accessible resources, thereby largely neglecting the variety of scrounging techniques individuals may use as well as the role of investment in food handling. Furthermore, although individual differences in boldness and social factors such as dominance have been described to influence foraging tactics, their potential interplay and effect in foraging contexts beyond the conventional producerescrounger game remains unclear. We investigated the relationship between social-foraging tactic use and dominance, pair bonds and boldness in a foraging experiment focused on food handling and alternative scrounging tactics. We conducted a producerescrounger experiment in a captive group of rooks in which individuals could produce by pulling up baited strings, or scrounge by retrieving fallen food items or joining a producer. There were three key findings: (1) dominant rooks adopted the producer tactic more often and more successfully than subordinates; (2) producing and scrounging by tolerance led to mixed benefits to paired birds; (3) bold birds scrounged by retrieving more often than shy birds. Importantly, individuals were highly consistent in their tactic use across conditions differing in food availability. Our study highlights the importance of taking both social factors and boldness (heterogeneity) into account when studying social-foraging dynamics and offers empirical data on food handling and alternative scrounging tactics that can be used to extend current models and experiments on social foraging.

Evidence suggesting that desire-state attribution may govern food sharing in Eurasian jays

State-attribution is the ability to ascribe to others an internal life like one’s own and to understand that internal, psychological states such as desire, hope, belief, and knowledge underlie others’ actions. Despite extensive research, comparative studies struggle to adequately integrate key factors of state-attribution that have been identified by evolutionary and developmental psychology as well as research on empathy. Here, we develop a behavioral paradigm to address these issues and investigate whether male Eurasian jays respond to the changing desire-state of their female partners when sharing food. We demonstrate that males feed their mates flexibly according to the female’s current food preference. Critically, we show that the males need to see what the female has previously eaten to know what food she will currently want. Consequently, the males’ sharing pattern was not simply a response to their mate’s behavior indicating her preference as to what he should share, nor was it a response to the males’ own desire-state. Our results raise the possibility that these birds may be capable of ascribing desire to their mates.

Caching at a distance: a cache protection strategy in Eurasian jays.

A fundamental question about the complexity of corvid social cognition is whether behaviours exhibited when caching in front of potential pilferers represent specific attempts to prevent cache loss (cache protection hypothesis) or whether they are by-products of other behaviours (by-product hypothesis). Here, we demonstrate that Eurasian jays preferentially cache at a distance when observed by conspecifics. This preference for a ‘far’ location could be either a by-product of a general preference for caching at that specific location regardless of the risk of cache loss or a by-product of a general preference to be far away from conspecifics due to low intra-species tolerance. Critically, we found that neither by-product account explains the jays’ behaviour: the preference for the ‘far’ location was not shown when caching in private or when eating in front of a conspecific. In line with the cache protection hypothesis we found that jays preferred the distant location only when caching in front of a conspecific. Thus, it seems likely that for Eurasian jays, caching at a distance from an observer is a specific cache protection strategy.

Inequity aversion in human adults: testing behavioural criteria from comparative cognition.

Inequity aversion refers to an attempt to reduce a perceived discrepancy between one’s own input and output ratio (i.e. ratio between work invested and rewards obtained) and that of others. It has been proposed that inequity aversion might also play a role in the decision-making process of other animals. One issue, however, is that while studies in comparative cognition define clear behavioural criteria for an inequity aversion effect in animals, studies conducted on humans rely on an implicit definition of the concept involving assumptions for which there is yet no experimental evidence. In particular, the basis of inequity aversion is assumed to be a social comparison process that is further motivated by fairness concerns. Based on the studies on non-human animals, we tested whether inequity aversion in adult humans will satisfy the behavioural criteria for these assumptions. Although humans showed a decrease in working for a lower-value reward when working alone (non-social effect), this effect was enhanced when a partner was present (social effect) suggesting that their inequity aversion might be based on a social comparison process. Additionally, our tests ruled out the possibility that selfish motives underlie the subjects’ decisions, which raises the possibility that fairness concerns might have been the primary motive. Our results thus show that human behaviour satisfies two criteria imposed by the definition of inequity aversion. A similar pattern is shown in other species, suggesting that the same or similar processes might influence economic decision-making in both humans and non-human animals.

Current desires of conspecific observers affect cache-protection strategies in California scrub-jays and Eurasian jays

Summary Many corvid species accurately remember the locations where they have seen others cache food, allowing them to pilfer these caches efficiently once the cachers have left the scene [1]. To protect their caches, corvids employ a suite of different cache-protection strategies that limit the observers’ visual or acoustic access to the cache site 2, 3. In cases where an observer’s sensory access cannot be reduced it has been suggested that cachers might be able to minimise the risk of pilfering if they avoid caching food the observer is most motivated to pilfer [4]. In the wild, corvids have been reported to pilfer others’ caches as soon as possible after the caching event [5], such that the cacher might benefit from adjusting its caching behaviour according to the observer’s current desire. In the current study, observers pilfered according to their current desire: they preferentially pilfered food that they were not sated on. Cachers adjusted their caching behaviour accordingly: they protected their caches by selectively caching food that observers were not motivated to pilfer. The same cache-protection behaviour was found when cachers could not see on which food the observers were sated. Thus, the cachers’ ability to respond to the observer’s desire might have been driven by the observer’s behaviour at the time of caching.

Are owners’ reports of their dogs’ ‘guilty look’ influenced by the dogs’action and evidence of the misdeed?

Highlights • We test cues that trigger dogs’ ‘guilty look’ behaviours.• The dogs’ action and evidence of the misdeed have no effect on the ‘guilty look’.• Dogs might not show the ‘guilty look’ in the absence of concurrent scolding by their owners.

Comparing the face inversion effect in crows and humans

Humans show impaired recognition of faces that are presented upside down, a phenomenon termed face inversion effect, which is thought to reflect the special relevance of faces for humans. Here, we investigated whether a phylogenetically distantly related avian species, the carrion crow, with similar socio-cognitive abilities to human and non-human primates, exhibits a face inversion effect. In a delayed matching-to-sample task, two crows had to differentiate profiles of crow faces as well as matched controls, presented both upright and inverted. Because crows can discriminate humans based on their faces, we also assessed the face inversion effect using human faces. Both crows performed better with crow faces than with human faces and performed worse when responding to inverted pictures in general compared to upright pictures. However, neither of the crows showed a face inversion effect. For comparative reasons, the tests were repeated with human subjects. As expected, humans showed a face-specific inversion effect. Therefore, we did not find any evidence that crows—like humans—process faces as a special visual stimulus. Instead, individual recognition in crows may be based on cues other than a conspecific’s facial profile, such as their body, or on processing of local features rather than holistic processing.

Difficulties when using video playback to investigate social cognition in California scrub-jays (Aphelocoma californica)

Previous research has suggested that videos can be used to experimentally manipulate social stimuli. In the present study, we used the California scrub-jays’ cache protection strategies to assess whether video playback can be used to simulate conspecifics in a social context. In both the lab and the field, scrub-jays are known to exhibit a range of behaviours to protect their caches from potential pilferage by a conspecific, for example by hiding food in locations out of the observer’s view or by re-caching previously made caches once the observer has left. Here, we presented scrub-jays with videos of a conspecific observer as well as two non-social conditions during a caching period and assessed whether they would cache out of the observer’s “view” (Experiment 1) or would re-cache their caches once the observer was no longer present (Experiment 2). In contrast to previous studies using live observers, the scrub-jays’ caching and re-caching behaviour was not influenced by whether the observer was present or absent. These findings suggest that there might be limitations in using video playback of social agents to mimic real-life situations when investigating corvid decision making.